Caitlin Elaine Mader
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Ecology and life history of Coccophagus gossypariae (Hymenoptera: Aphelinidae), a parasitoid of Eriococcus spurius (Hemiptera: Eriococcidae) by Caitlin Elaine Mader A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science In Forest Biology and Management Department of Renewable Resources University of Alberta © Caitlin Elaine Mader, 2018 Abstract The American elm (Ulmus americana) is a valuable component of urban forests in Alberta. In many Alberta municipalities, the health of these trees is being heavily impacted by the invasive scale insect Eriococcus spurius (Hemiptera: Eriococcidae). Biological control of scales using hymenopteran parasitoids can be an effective replacement or addition to control using chemical insecticides. However, no effective biological control agent has been found for E. spurius. This project investigates the life history and host interactions of Coccophagus gossypariae (Hymenoptera: Aphelinidae), a parasitoid wasp that was discovered parasitizing E. spurius in Calgary. Field studies took place at 46 sites in Calgary during 2015, and 44 of the same sites in 2016. At each site, I conducted repeated-measures sampling every two weeks for eight rounds in 2015, and 5 rounds in 2016. At each site and sampling round, I collected three types of samples: Adult C. gossypariae that had emerged from their hosts and had been caught in mesh traps over the preceding two weeks; first instar E. spurius nymphs that had been caught on sticky traps over the preceding two weeks; and live adult scale insects, which I later dissected in the laboratory for fecundity and parasitism data. I used various combinations of the resulting datasets, along with climate data and urban forest inventory data supplied by the City of Calgary to answer different questions in three investigative chapters. The chapters 2-4 each provide a different area of focus in better understanding this tri-trophic system. In the first chapter, I undertook basic investigations of C. gossypariae life history traits. I found that it is well established throughout Calgary, and accounts for 98% of the parasitism of E. spurius. Parasitism rates varied greatly between sites, ranging from 6.8% to 81.0%. I recorded a sex ratio of 7% males, and a strong avoidance of superparasitism. In the second chapter, I adopted a much broader focus, conducting spatial analyses on how urban landscape factors influence E. spurius populations. I found that the amount of impermeable surfaces within 10 m from a tree, and the number of other elm trees ii upwind of it are both positively associated with higher E. spurius densities. Finally, I examine how E. spurius and C. gossypariae interact over the course of the season, comparing their relative phenologies, and the effects of parasitism on E. spurius reproduction. I found that E. spurius have already finished reproduction before the point in the season when the majority of parasitism-induced mortality occurs. Because E. spurius adults will not live to reproduce again, most mortality induced by C. gossypariae emergence after reproduction will have little impact on its host’s population. However, E. spurius that are parasitized by C. gossypariae produce significantly fewer eggs than those that are not. I conclude that if C. gossypariae has any suppressive effects on its host`s population, it is likely due to fecundity effects, rather than mortality effects. I did not, however, document any influence of C. gossypariae parasitism on changes in E. spurius population. This study provides some first steps for future work on whether C. gossypariae can be used in biological control of E. spurius. iii Acknowledgements This project would not have been possible without the funding provided by the City of Calgary`s Urban Conservation Unit, the Alberta Conservation Association`s Grants in Biodiversity, and the Anina Hundsdoerfer Memorial Graduate Scholarship. Thanks to Dr. Nadir Erbilgin for the tremendous scientific and logistical support, Jim Watts for supporting this project from the very beginning, Lien Luong and John Acorn for serving as my defense committee, Zhong Li and Todd Kemper for the kind references that got me here and helped with funding, Susan Ryan, John Mader, and Emily Upham-Mills for editing help, my family, Amélie Roberto-Charron, Kelli Luber, Chris Parsonage, Meghan Schech, and Patrick Chan for moral support when it was needed most, the extended Edmonton circus community, Linnea’s Legion, and all of my house- mates for keeping me sane, Daniel Gorham, Pranamika Dutta, April Papequash, and Ashley Pisesky for your care and attention to thousands of incredibly tiny samples, Erik Luber for answering all of my math questions, especially the ones I never ended up using, Jonathan Cale for answering all of my stats and R questions, Evelyn Merrill for guidance in spatial analyses, Violet Zhao, Sanat Kanekar and Jen Klutsch for making the office so enjoyable, and everyone in the Erbilgin lab for their feedback and camaraderie. iv Contents Abstract ........................................................................................................................................... ii Acknowledgements ........................................................................................................................ iv Chapter 1: Thesis Introduction........................................................................................................ 1 Tables and Figures ...................................................................................................................... 5 Chapter 2: Life history traits of Coccophagus gossypariae (Hymenoptera: Aphelinidae) in field studies ............................................................................................................................................. 7 Abstract ....................................................................................................................................... 7 Introduction ................................................................................................................................. 8 Materials and Methods .............................................................................................................. 13 Sampling scheme .................................................................................................................. 13 Tree selection ........................................................................................................................ 13 Sampling Eriococcus spurius and parasitoid specimens ...................................................... 14 Sample Processing ................................................................................................................ 16 Descriptive statistics ............................................................................................................. 16 Analysis of superparasitism .................................................................................................. 17 Analysis of prolonged effects of imidacloprid on percent parasitism .................................. 17 Results ....................................................................................................................................... 18 Parasitoid emergence rates and species assemblage ............................................................. 18 Coccophagus gossypariae sex ratios .................................................................................... 19 Parasitism and superparasitism rates .................................................................................... 19 Effect of imidacloprid on hymenopteran parasitism ............................................................. 20 Anecdotal field observations................................................................................................. 20 Discussion ................................................................................................................................. 21 Parasitoid species assemblage and geographic distributions ................................................ 21 Coccophagus gossypariae sex ratios .................................................................................... 23 Long-term imidacloprid effects on parasitoids ..................................................................... 23 Rates of parasitism and superparasitism ............................................................................... 24 Anecdotal field observations................................................................................................. 26 Future directions ................................................................................................................... 27 Tables and Figures .................................................................................................................... 29 v Chapter 3: Spatial and environmental predictors of ..................................................................... 36 post-insecticide Eriococcus spurius recolonization of urban elm trees ........................................ 36 Abstract ..................................................................................................................................... 36 Introduction ............................................................................................................................... 36 Materials and Methods .............................................................................................................